Oviposition Behavior of an Aberrant African Stingless Bee Meliponula Bocandei, with Notes on the Mechanism and Evolution of Oviposition Behavior in Stingless Bees 1)
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Oviposition Behavior of an Aberrant African Stingless Bee Meliponula bocandei, with Notes on the Mechanism and Title Evolution of Oviposition Behavior in Stingless Bees (With 16 Text-figures and 6 Tables) Author(s) SAKAGAMI, Shôichi F.; ZUCCHI, Ronaldo; ARAÚJO, Virgílio de Portugal Citation 北海道大學理學部紀要, 20(4), 647-690 Issue Date 1977-05 Doc URL http://hdl.handle.net/2115/27625 Type bulletin (article) File Information 20(4)_P647-690.pdf Instructions for use Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP Oviposition Behavior of an Aberrant African Stingless Bee Meliponula bocandei, with Notes on the Mechanism and Evolution of Oviposition Behavior in Stingless Bees 1) By Shoichi F. Sakagami, Ronaldo Zucchi and Virgilio de Portugal Araujo Zoological Institute, Hokkaido University, Sapporo, Departamento de Biologia, Faculdade de Filosofia, Ciencias e Letras de Ribierll.o Preto, Universidade de Sll.o Paulo, Ribeirlio Preto and Instituto Nacional de Pesquisas da Amazonia, Manaus (With 16 Text-figures and 6 Tables) Meliponula bocandei (Spinola 1853) is a singular African stingless bee, showing morphological features intermediate between Melipona and the other stingless bees (Wille 1963)_ As a continuation of our serial reports on the oviposition behavior of stingless bees, the present paper deals with behavioral observations of this species, only a summary of which was previously reported (Sakagami and Zucchi 1966). In this occasion, our previous hypothesis on the mechanism underlying the oviposition process of stingless bees will be revised, accompanied with some discussions on related problems_ It was unfortunate that this species, highly aberrant in behavior from other stingless bees, was studied at an earlier stage of our serial observations, when our comparative knowledge was inevitably imperfect. But the basic behavioral characters given below will be useful for further detailed studies. In subsequent pages the other species observed by us and cited for comparisons will be shown, unless necessary, with generic or subgeneric names alone, according to the system by Moure (1951, 1961) adopted in our serial reports. (Other systems were recently presented by Wille and Michener 1973, and partly by Sakagami 1975). The species names, authors and full citations of the behavioral accounts are found in the references at the end of the paper. The generic and subgeneric names not found there are those cited from unpublished data of ours or our students. For simplicity these taxa are henceforth regarded as genera. The terminology on behavioral details revised by Sakagami and Zucchi (1974) will be adopted in subsequent descriptions. 1) Behavior studies of the stingless bees, with special reference to the oviposition process. X. Jour. Fac. Sci. Hokkaido Univ. Ser. VI, Zool. 20 (4), 1977. 647 648 S. F. Sakagami, R. Zucchi and V. de P. Araujo The observed colony was introduced from Luanda, Angola, to Brazil, and had been kept in the apiary of Departamento de Biologia, Faculdade de Filosofia, CiE'llcias e Letras de Rio Claro, the State of Silo Paulo, in a hive following the pattern developed by Araujo (1956). For the behavior study, the colony was divided and the old queen accompanied with brood, food and adult workers was transferred in an observation hive (Sakagami 1966) and observed mainly from December 26, 1962 to February 28, 1963. Meliponula is a very productive bee. Its nests often become fairly large, the extent of brood area occasionally attaining 15",18 cm in diameter and 20", 25 cm high (Araujo (1955). It is posRible that the behavior in the small colony kept in an observation hive deviates in some aspects from that in natural nests. But from observations of oviposition behavior in Melipona rufiventris Lepeletier under diverse colony conditions, Camilo-Atique (1974) clarified the persistence of qualitative aspects of behavior patterns and their articulation, though quantitative aspects varied according to colony conditions. This would be valid for Meliponula, too. 1. Behavior out of oviposition process 1.1. General and miscellaneous accounts Meliponula builds incomplete combs (Fig. 1), the type being relatively rare in stingless bees, known only in some limited genera, Friesella, Duckeola, etc. Cells are juxtaposed one another but not neatly lie in the same plane. Each cell retains spherical contour at the top and the bottom, exhibiting an uneven appearance of comb surface. Combs expand irregularly, not concentrically as in typical comb builders and are occasionally but not always surrounded with involucrum. In general appearance at; well a" in size and coloration, combs are similar to those of Duckeola (Kerr et al. 1966). In :VIarch 1963, a piece of Duckeola comb, ca. 3 X 4 cm sq., waR given to the nest. It was destroyed within a day. A piece of honeybee comb with stored honey was accepted and covered with cerumen, but destroyed after honey consumption. Although no detailed observation on intranidal worker behavior was made, no particular difference from other genera was noted. General motor pattern Fig. 1. Combs of 1l1eliponula bocandei. C = Cells in construction, R = Royal cell. Oviposition behavior of Meliponula 649 resembles that of Melipona, not so sluggish as in Duckeola but slower than in most other genera. At walking wings are not rhythmically fluttered and food regurgitation between workers is often made with a twisted posture as in other genera. Feces and other debriR are accumulated on the floor of the hive, forming several heaps, which are actively carried away on the wing during daytime. For this active removal the colony did not suffer the damage by the phorid fly, Pseudohypocera kerteszi (Enderlein), the worst enemy of stingless bees in southern Brazil, depsite many flieR invaded the colony and workers did not pay much attention to them. Newborn ,yorkers were mostly helped by older sisters at emergence. As in other genera, at ventilation the head iH directed against, not to the nest entrance. Usually four to five workerfi stay at nest entrance aH guards. They exhibit spontaneous repetition of fore- and backward movements, but never so overtly as in Melipona qU(ldrifasciata. Guards seldom examined the homing foragers, responding with a mild retreat. When interfered they first retreat. By continuous stimulation with a stick, some guards advance, raising the fore body and fore legs and opening mandibles, dart and gnaw the stick. No further aggression nor ritualized defensive posture Ruch as the warning fanning (Scaptotrigona, Melipona seminigra), etc. are exhibited. From pigmentation i" concluded that most Fig. 2. A young, inseminated queen (A, C) and a foraging worker (E, D) of Meliponula bocandei. A, B = Dorsal views (\Vorker = Dried specimen. In fresh condition, the metasoma is longer). C, D=Head viewed frontally. i:lcale = 1 mm for A, B; 0.5 mm for C, D. 650 S. F. Sakagarni, R. Zucchi and V. de P. Araujo guards are at ages intermediate between house and foraging bees. Once a pollen laden worker participated in guarding for more than one hour. 1.2. Queen behavior The queen and worker are shown in Fig. 2 and differences in some metric characters in Table 1. The head of the queen is a triffie narrower than in workers, but the mesosoma is slightly wider. The queen walks more rapidly than in Melipona and Scaptotrigona. Wings are beaten rather sporadically at walking but rhythmically at resting. Beats are single-stroked without vibration. The time spent for ten strokes are 4.0 sec (l case), 4.5 (3), 5.0 (2), 6.0 (6), 6.5 (3), 7.0 (5), 9.0 (2) and 10.0 (1). The queen behavior is characterized by two features which are only rarely observed in other genera. One is the violent and constant shaking of antennae distinctly longer than in workers (Fig. 2, Table 1), irrespective of walking or resting, which is only similar to the behavior presented by the queen of Axestotrigona, another African group. The other is the incessant crUlsmg throughout the hive, not only on the floor, combs and storage pots but even beneath the glass lid. Moreover, she frequently visited the entrance corridor connecting the hive to outdoors, even reaching the nest entrance. This was first Table 1. Size difference between queen (n=l) and worker (n=lO) in some metric characters (in mm) arranged in the ascending order of queen/worker ratio (L=length, W =width). Worker Queen/worker Character Queen Mean SD ratio Eye L 1.556 1.926 0.033 0.808 Hind basitarsus W 0.629 0.744 0.047 0.846 Mesoscutellum L 0.741 0.874 0.065 0.847 Hind tibia W 0.963 1.133 {).047 0.850 Head L 2.370 2.659 0.119 0.891 Head \V 3.037 3.356 0.067 0.905 Fore wing L 6.444 6. 763 0.094 0.953 Mid tibia L 1.852 1.926 0.047 0.962 Hind wing L 4.963 5.089 0.105 0.975 Mesoscutum L 2.000 2.037 0.076 0.982 Mesosoma \V 3.926 3.474 0.102 1.130 Mid femur L 2.148 1.896 0.068 1.133 Hind basitarsus L 1. 556 1.363 0.089 1.141 Scape L 1.444 1.230 0.049 1.175 Hind tibia L 2.815 2.363 0.070 1.191 Hind femur L 2.630 2. 104 0.076 1.250 Mid basitarsus L 1. 704 1.318 0.055 1. 292 Pedicel + Flagella L 3.286 2.428 0.055 1.3540 Flagellomere I L 0.370 0.258 0.021 1. 437 Malar L 0.222 0.151 0.059 1.470 Flagellomere II L 0.333 0.221 0.004 1.lil0 Oviposition behavior of Meliponula 651 suspected as an instability caused by some abnormal colony conditions.